Peckhamia 171.1 Oophagy by semicupreus 1

PECKHAMIA 171.1, 12 September 2018, 1―6 ISSN 2161―8526 (print) urn:lsid:zoobank.org:pub:95DB4AA8-9498-418F-8BB1-972096BDA1E2 (registered 11 SEP 2018) ISSN 1944―8120 (online)

Oophagy by (Araneae: Salticidae: Plexippina)

Javed Ahmed 1, David E. Hill 2, Richard J. Pearce 3, A. N. Suresh Kumar 4, Rajashree Khalap 5 and Krishna Mohan 6

1 Panchavati Housing Society, Building No. A/3, Flat No. H/8; Opp. Police Camp, Vijay Nagar, Marol Maroshi Road, Andheri (East), Mumbai 4000, , email [email protected] 2 213 Wild Horse Creek Drive, Simpsonville, SC 29680-6513, USA, email [email protected] 3 Moreton Morrell College, Warwickshire, UK, email [email protected] 4 Nataraj Palace, First Floor, New #54, Saravanan Street, T. Nagar, Chennai, Tamil Nadu 600017, India, email [email protected] 5 5-A, Sagar Sangeet, 58 Shahid Bhagat Singh Marg, Colaba, Mumbai 400005, India, email [email protected] 6 Prabhu Hospital, Hospital Cross Road, Moodubidire 574227, India, email [email protected]

Abstract: A female Hyllus semicupreus was observed feeding on the eggs of a coreid hemipteran in Andhra Pradesh, India.

Key words: Anastatus, Andhra Pradesh, biocontrol, Coreidae, egg feeding, jumping

Hyllus C. L. Koch, 1846 is a large tropical Afroeurasian that includes mid-sized to large jumping with stout, hirsute bodies, a rounded carapace, and in many species tufts of long setae on either side of the eye region (Wesołowska, 2008; Caleb et al. 2014). Of the 71 species presently placed in this genus only five have been reported from India (Table 1; WSC 2018).

Table 1. Jumping spiders of the genus Hyllus reported from India. species distribution notes H. bos (Sundevall, 1833) India Named in a short note without a description. Simon (1903) guessed that this species probably belongs in Hyllus. H. diardi (Walckenaer, 1837) widespread from south to southeast Asia This is the best-known member of the genus, and Indonesia (Sunda) frequent in the international pet trade. H. manu (Caleb, Christudhas, Chennai, South India Known only from the Madras Christian College Laltanpuii & Chitra, 2014) campus. H. pudicus (Thorell, 1895) Tharawaddy, and India According to Prószyński (1992) distribution in south to southeast Asia in uncertain. H. semicupreus (Simon, 1885) India and Sri Lanka according to published One of the better-known Hyllus that may also be accounts, but photographed in Malaysia common in southeast Asia and Sunda.

The natural history, including behaviour, biology and distribution of these spiders, with few exceptions (Ahmed & Satam, 2015; Gilman, 2016), is little known. In this paper we document, for the first time, oophagy by a female Hyllus semicupreus (Figure 1), a widely-distributed and relatively well-known species in India and Sri Lanka that may also be found far to the southeast from the Malayan peninsula to Borneo (Figure 2). Peckhamia 171.1 Oophagy by Hyllus semicupreus 2

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12 15 6 Figure 1. Oophagy by a female Hyllus semicupreus in Andhra Pradesh. 1-2, Feeding on the first egg in a line of 16 coreid eggs. 3, Subsequent feeding on the second egg in this line. 4-6, Line of eggs after this spider finished feeding. Only 7 (numbered) of the 16 eggs were eaten. Note the small eupelmid (cf. Anastatus sp.) wasp (5-6). Photographs by A. N. Suresh Kumar. Peckhamia 171.1 Oophagy by Hyllus semicupreus 3

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1- Simon 1885 2- Tikader 1974 3- Tikader & Biswas 1981 4- Sharma et al. 2013 5- Prószyński 2014 6- Ahmed & Satam 2015 7- Roy et al. 2016 8- internet photographs 9- this paper

Figure 1. Range of Hyllus semicupreus in south and southeast Asia. Although published records for this species are confined to India and Sri Lanka, photographs posted on the internet (8) indicate that it is widely distributed from Delhi to Borneo. The background relief map is in the public domain (CC0), courtesy of maps-for-free (https://maps-for-free.com).

This incident of oophagy (Figure 1) was documented by one of the authors (ANSK) while photographing terrestrial invertebrates on privately owned agricultural land, located at Pellakuru Mandal, Nellore District, Andhra Pradesh, India (25 AUG 2018, 09:00, N13.8305399°, E79.8096502°). Initially this author approached a lone dragonfly perched atop a dried stem, part of a natural barricade system of thorny shrubbery bordering the property to dissuade cattle and intruders alike from trespassing, a common practice in the countryside across India. Upon drawing near, a female Hyllus semicupreus was observed resting on a row of bright red eggs. Based on their distinctive shape and coloration we have subsequently determined that these eggs had been laid by a leaf-footed bug (Hemiptera: Coreidae). This spider remained absolutely motionless as she fed, even when approached and photographed from various directions. In order to get a better look at the egg under the spider, the spider was gently nudged, and as she proceeded to move further along the row of eggs, the egg she was sitting on was found to be cut open and the contents consumed. The spider was then left in place. Peckhamia 171.1 Oophagy by Hyllus semicupreus 4

When the site was revisited later in the day (~16:30), this spider was missing. The row of eggs was then photographed (Figure 1:4-6). Out of a total of 16 eggs that had been deposited on the stem, ranging in length from ~1.95-2.05 mm, seven had been consumed, while the remaining nine eggs were still intact. At this time the eggs were also visited by a diminutive parasitoid wasp (Hymenoptera: Eupelmidae, cf. Anastatus sp.; Figure 1:5-6). After inspecting the eggs for several seconds this wasp flew away, leaving the intact eggs that remained untouched. Wasps of the genus Anastatus are known to parasitise the eggs of a wide range of insects, including Blattaria, Hemiptera, Lepidoptera, Mantodea, Neuroptera, Orthoptera, Heteroptera and Phasmida (Hanson & Gauld, 1995).

Oophagy by salticid spiders. Oophagy (egg eating) by salticid spiders may be relatively common and this behavior has now been documented in a number of species (Table 2).

Table 2. Records of oophagy by salticid spiders. Clade is based on Maddison (2015). species clade references notes sp. Spartaeina Jackson & Hallas, 1986 Fed on eggs of theridiid and amaurobiid spiders. Cosmophasis cf. Chrysillini Donovan & Hill, 2017 Female fed on a series of butterfly (Lepidoptera: thalassina Nymphalidae) eggs. Cyrba sp. Spartaeina Jackson & Hallas, 1986; Fed on eggs of theridiids and amaurobiids. In Hallas, 1988; Jackson, captivity some hatchlings fed on eggs in brood sac. 1990 Evarcha albaria Plexippina Cushing, 2012 Robbed ants of prey and brood (eggs and larvae). coecatus Harmochirina Nyffeler et al., 1990 Fed on insect eggs. sp. Dendryphantina Nyffeler et al., 1990 Fed on insect eggs. Dendryphantina Pfannenstiel, 2008 Fed on eggs of Lepidoptera. Hyllus semicupreus Plexippina this study Female fed on eggs of a coreid bug. Jacksonoides Astiini Jackson, 1988a Fed on unattended eggs of other salticid spiders. queenslandica Menemerus fulvus Chrysillini Cushing, 2012 Robbed ants of prey and brood (eggs and larvae). Metaphidippus flavipedes Dendryphantina Nyffeler et al., 1990 Fed on insect eggs. Metaphidippus galathea Dendryphantina Nyffeler et al., 1990 Fed on insect eggs. Neobrettus cf. tibialis Spartaeina Ahmed et al., 2018 Females fed on their own and conspecific eggs, and may feed on eggs of other salticids. Opisthoncus necator Viciriini Nyffeler et al., 1990 Fed on spider eggs. Paracyrba wanlessi Spartaeina Żabka & Kovac, 1996 Fed on insect eggs. sp. Dendryphantina Nyffeler et al., 1990 Fed on insect eggs. Phidippus audax Dendryphantina Nyffeler et al., 1990 Fed on insect eggs. Phyaces comosus ? Simaethina Jackson, 1986b; Moffet, Fed on eggs of larger salticids. 1991 Plexippus setipes Plexippina Cushing, 2012 Robbed ants of prey and brood (eggs and larvae). Portia sp. Spartaeina Jackson & Hallas, 1986; Fed on eggs of other spiders. Hallas, 1988 Portia fimbriata Spartaeina Jackson & Blest, 1982; Fed on eggs of other spiders. Jackson, 1985, 1986b Siler cupreus Chrysillini Cushing, 2012 Robbed ants of prey and brood (eggs and larvae). Tauala lepidus Astiini Jackson, 1988b Fed on eggs of conspecifics and other salticids. Peckhamia 171.1 Oophagy by Hyllus semicupreus 5

Ecological significance. Spiders are significant terrestrial predators, consuming an estimated 400-800 million tons of prey annually (Nyffeler & Birkhofer, 2017). Although parasitoids such as the eupelmid wasps (e.g. Anastatus spp.) are considered to be potentially important agents for the biocontrol of harmful insects (Roversi, 1990; Ulber et al., 2010; Ramos et al., 2018), the potential for biocontrol by spiders has been open to more debate (Betz & Tscharntke, 2017; Cotes et al., 2018; Ndava et al., 2018). In part this is due to the notion that spiders are generalists and in part due to the assumption that prey must be moving in order to attract the attention of spiders. Given the importance of eggs and larval stages in the life cycles of many invertebrates, the potential of spiders as biocontrol agents has sometimes been brought into question (Finke & Denno, 2004; Hodge, 1999). Increasing evidence shows that, whilst many spiders may play a generalist role, others are more selective in their feeding preferences. Consumption of eggs may have a greater impact on populations of coreid bugs than consumption of nymphs or adults which may already have caused plant damage. Hyllus semicupreus and eupelmid parasitoids like Anastatus may compete as important control agents for coreid bug populations. The relative impact of oophagous spiders like H. semicupreus and parasitoid wasps on populations of coreid bugs and other insects deserves further study.

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